[Hexapoda phylogeny: implications for an aquatic origin hypothesis]

Emergence from an aquatic environment to the land is one of the major evolutionary transitions within the arthropods. It is often considered that the first hexapods, and especially the first Collembola, went from the sea through intermediate freshwater environments to colonize fully terrestrial ecosystems. To understand the ancestral ecology of hexapods, a phylogenetic framework is used. By mapping ecological attributes onto the phylogeny, it is shown that hexapods colonized terrestrial environments directly from marine environment without a transition through freshwater. An edaphic life-style is the basal state for Collembola and more generally for hexapods as a whole. Aquatic ecology is inferred to be a secondary change that occurred several times independently, particularly in some group of Collembola and Pterygota. The answer is ambiguous for Pterygota, whether the first Pterygota had aquatic larvae and reversed toward fully terrestrial in Neoptera, or aquatic larvae appeared independently in Odonata and Ephemeroptera. Subsequently, aquatic larvae were secondarily acquired in various groups independently (e.g. Plecoptera, Trichoptera, Coleoptera).